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Amitava Rakshit • Harikesh Bahadur Singh
Editors
Advances in Seed Priming
Editors
Amitava Rakshit
Department of Soil Science and
Agricultural Chemistry
Institute of Agricultural Sciences, BHU
Varanasi, Uttar Pradesh, India
Harikesh Bahadur Singh
Department of Mycology and
Plant Pathology
Institute of Agricultural Sciences, BHU
Varanasi, Uttar Pradesh, India
ISBN 978-981-13-0031-8 ISBN 978-981-13-0032-5 (eBook)
https://doi.org/10.1007/978-981-13-0032-5
Library of Congress Control Number: 2018944402
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209
© Springer Nature Singapore Pte Ltd. 2018
A. Rakshit, H. B. Singh (eds.), Advances in Seed Priming,
https://doi.org/10.1007/978-981-13-0032-5_12
N. Ozbay (*)
Department of Horticulture, Agricultural Faculty, Bingol University, Bingol, Turkey
12
Studies onSeed Priming inPepper
(Capsicum annuum L.)
NusretOzbay
Abstract
Plants are regularly exposed to various adverse environmental conditions such as
drought, salinity, chilling, and high temperatures. These abiotic stresses adversely
affect the plant growth and productivity and, in extreme cases, cause plant death.
One feature expected of high-quality seeds is to be able to germinate under
adverse growing conditions. Rapid germination and emergence are essential for
successful crop establishment, for which seed priming could play an important
role. Seed priming is an effective technology to enhance rapid and uniform emer-
gence and to achieve high vigor, leading to better stand establishment and yield.
Over the past 50 years, there has been extensive literature published on seed
priming and related pre-sowing seed treatments. These have been particularly
promising for high-value small-seeded vegetable crops such as pepper where
rapid, uniform germination is at a premium either in the cell transplant produc-
tion or where accurate and uniform plant population density is required from
direct sowing. The aim of this chapter is to review some of the current ideas and
recent work on physiological priming treatments designed to enhance germina-
tion and emergence performances of pepper seeds.
Keywords
Pepper · Seed priming · Germination · Emergence · Seedling growth and estab-
lishment · Vigor · Environmental stress
210
12.1 Introduction
Pepper is one of the most important vegetable crops in the world. Peppers are grown
worldwide because of their adaptation to different agroclimatic regions and their
wide variety of shapes, sizes, colors, and pungencies of the fruit (Qin etal. 2014).
Bell pepper or sweet pepper belongs to the genus Capsicum, a member of the
Solanaceae family that includes tomatoes, potatoes, and eggplants. Pepper is a cul-
tivar group of the species Capsicum annuum L.Cultivars of the plant produce fruits
in different colors, including red, yellow, and orange, but more exotic colors include
purple, white, and lime green. The fruit is also frequently consumed in its unripe
form, when the fruit is still green. In the United States, Canada, the United Kingdom,
Ireland, and some other parts of the world, in addition to the terms “bell pepper” and
“sweet pepper,” the fruit is often referred to simply as a “pepper,” whereas in many
Commonwealth of Nations countries, such as Australia, India, Malaysia, and New
Zealand, they are called “capsicum.”
Peppers are native to Mexico, Central America, and northern South America.
Pepper seeds were later carried to Spain in 1493 and from there spread to other
European, African, and Asian countries (Andrews 1993; Abdel-Kader and El-Mougy
2014). According to FAO statistical database, in 2014, pepper is cultivated on
3,625,452ha in the world, with an annual production of 36,143,113 metric tons
(MT) representing an annual yield of 9.97MT ha−1. Today, China is the world’s
largest pepper producer, followed by Mexico, Turkey, and Indonesia (FAO 2017).
Peppers grow better in evenly moist and evenly warm soil and can be grown year
round in frost-free conditions. If moisture levels or temperature levels uctuate, too
much growers will have problems related to some diseases and yield. It has also
been reported that pepper seed germination and emergence are slow and nonuni-
form under normal as well as stressful conditions (Belletti and Quagliotti 1988;
Chartzoulakis and Klapaki 2000; Demir and Okcu 2004; Khan etal. 2009a). Low
ability of seed germination during seed production and rapid deterioration during
storage of pepper seed are also the major problems that affect seed quality (Siri etal.
2013). Rapid germination and emergence are essential for successful crop establish-
ment. Various seed treatments have been suggested to improve seed germination
and emergence, and one of them is priming. Seed priming is an effective technology
to enhance rapid and uniform emergence and to achieve high vigor, leading to better
stand establishment and yield (Singh etal. 2015; Ibrahim 2016).
During priming, the seeds are partially hydrated in such a way that pre-
germination metabolic activities start; however, radicle protrusion is prevented fol-
lowed by drying of seeds to the original moisture content (Sharma etal. 2015). Seed
is primed to obtain faster and more uniform germination resulting in a stronger crop
stand. Yadav etal. (2011) reported that priming treatments improved germination
percentage as well as rate of seed germination in pepper under normal as well as the
stressed conditions. Maiti etal. (2009) studied the effect of priming on seedling
vigor and productivity of pepper, during post-rainy seasons demonstrating that
N. Ozbay
211
priming improved germination and seedling development and yield pepper. Another
primary benet of priming is the extension of the temperature range at which a seed
can germinate (Valdes and Bradford 1987; Ellis and Butcher 1988). This has par-
ticular importance for crops such as pepper, in which seedlings are produced in the
early spring at low temperatures for planting in the open eld and in late summer at
higher temperatures for glasshouse production. It has been reported that priming
improved seed germination at suboptimal temperatures (Pandita et al. 2007;
Korkmaz and Korkmaz 2009). Nascimento et al. (2011) reported that priming
improved the physiological quality of hot pepper seeds by improving germination at
high temperature (35°C).
12.2 Seed Priming Methods
In seed technology, different pre-sowing treatments have been utilized to reduce the
time from sowing until germination or emergence and to shorten the interval
between the rst and last seed germination. One of the most important pre-sowing
treatments is known as seed priming. The principle of priming is based on the fact
that it is possible to hydrate seed in some ways at a moisture level sufcient to initi-
ate the early events of germination but not sufcient to permit radical protrusion
through the seed coat (Sivritepe and Sivritepe 2016). Seed priming enhances seed
performance by rapid and uniform germination, more uniform emergence, normal
and vigorous seedlings, greater tolerance to environmental stress, and reduced dor-
mancy in many species (Khan 1992; Desai etal. 1997; Cantliffe 2003). Additionally,
this physiological treatment provides invigoration treatment to partially aged seed
lots. Excellent review studies have considered the importance of seed priming and
the factors that affect its success (Khan 1992; Parera and Cantliffe 1994; Bray 1995;
Pill 1995; Taylor et al. 1998; Welbaum etal. 1998; McDonald 2000). Benecial
effects from priming have been reported for several vegetable seeds including pep-
per (Gomes etal. 2012). Improvement of germination in pepper plant by priming
with water and NaCl has been reported (Smith and Cobb 1991a). The results of
Kaya etal. (2010) showed that priming pepper seeds can increase germination per-
centage and accelerate germination at suboptimal temperatures. Priming pepper
seeds in PEG and KNO3 solutions signicantly increased emergence percentage and
decreased mean time to emergence of pepper seedlings (El-Shatoury 2010). Various
priming treatments have been developed to enhance seed germination, seedling
growth, and yield in most of the crops under normal and stress conditions and also
to avoid biotic and abiotic stresses during germination and emergence phases
(Bradford 1986; McDonald 2000). These priming methods can be listed as follows
(Ashraf and Foolad 2005; Lutts et al. 2016; Sivritepe and Sivritepe 2016):
hydropriming, osmopriming, solid matrix priming, halopriming, drum priming,
hormopriming, thermopriming, biopriming, nutripriming, and organic priming.
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
212
12.2.1 Hydropriming
It is a simple and low-cost priming technique in which seeds are soaked in water
(aerated distilled water is preferred) for a certain period of time to a point where
pre-germination metabolic activities start without actual germination and subse-
quently dried back to storage moisture contents before sowing (Singh etal. 2015).
This process is especially useful in economically disadvantaged, arid crop-growing
areas. Hydropriming has been considered as a simple and cost-effective strategy for
improving germination and emergence of pepper. For example, Demir and Okcu
(2004) reported that aerated hydration treatment of pepper and eggplant seeds sig-
nicantly increased nal germination of both species over a wide range of tempera-
ture (18–35 °C). Hydropriming with normal water or warm water improved the
radicle protrusion of pepper seeds signicantly as compared to the control treatment
(Yadav etal. 2011). Similarly, hydropriming of pepper seeds for 24h enhanced the
germination and emergence percentage, ensured early germination and seedling
emergence, increased uniformity of emergence, and increased seedling vigor com-
pared to unprimed seeds (Uche etal. 2016).
12.2.2 Osmotic Priming
This method is originally developed and described by Heydecker etal. (1973, 1975).
Osmotic priming technique, also called osmopriming or osmoconditioning, is based
on the controlled hydration of seeds to a level that allows pre-germination metabolic
activity but inhibits radicular emergence. It is achieved by immersing seeds in an
aqueous solution of a chemically inert but osmotically active compound such as
polyethylene glycol (PEG) for a specic period at a specic temperature (Heydecker
etal. 1975; Khan 1992). At the end of the process, seeds are rinsed before further
processing. Taylor and Harman (1990) and Gray (1994) provide access to the very
large literature on osmotic priming. Some of the compounds that are used for
osmotic priming include sugars, polyethylene glycol (PEG), glycerol, sorbitol, or
mannitol. The improvement of germination and emergence performance of seeds of
many plant species through a pre-sowing treatment with a chemically inert osmoti-
cum have been reported by many researchers (Bennett etal. 1992). The effect of
seed priming on eld emergence of pepper seeds has ranged from no improvement
to some advancement in mean germination time (Yaklitch and Orzolek 1977;
Bradford etal. 1990). Aljaro and Wyneken (1985) reported that osmoconditioning
of sweet pepper seed did not affect the germination percentage but shortened germi-
nation time and gave more uniform germination. Pepper seeds primed in mannitol
had improved nal percent germination and produced larger seedlings than
nonprimed seeds (Georghiou etal. 1987; Passam etal. 1989). Kikuti et al. (2005)
reported that sweet pepper seeds primed with PEG 6000 performed better in all the
vigor tests assessed, except the germination test. Demirkaya (2006) studied osmotic
conditioning with polyethylene glycol to enhance the germination percentage and to
N. Ozbay
213
reduce the mean germination time of pepper seeds. He found that osmoconditioned
seed enhanced germination percentage and reduced the mean germination time.
Cortez-Baheza etal. (2007) indicate osmopriming treatments using PEG in combi-
nation with GA3 or KNO3 have shown to be good approaches to revigorate pepper
seeds for commercial purposes. Siri etal. (2013) reported that seed germination was
improved in osmoprimed pepper seeds primed in a PEG 6000 solution with the
osmotic potential of −1.5MPa for 6days.
12.2.3 Halopriming
In halopriming, the seeds are immersed in different inorganic salt solutions (NaCl,
KNO3 CaCl2, CaSO4, etc.) which facilitate the process of seed germination and
subsequent seedling emergence even under adverse environmental conditions.
Halopriming is a simple and cheap agro-technique and therefore found suitable to
be recommended to the farmers owing to better synchrony of emergence and crop
stand under various conditions of environment (Sedghi etal. 2010). A number of
studies have shown a signicant improvement in seed germination, seedling emer-
gence and establishment, and nal crop yield under unfavorable conditions such as
suboptimal temperature and salinity as a result of seed halopriming. For example,
Smith and Cobb (1991a) reported that priming with water and NaCl improved ger-
mination in pepper plant. Carter (1994) initiated a work to examine the effects of
NaCl priming on total germination and the rate of germination under suboptimal
temperature conditions. Seeds of “Tam Veracruz” pepper were primed at 23°C for
5days in double-distilled H2O, or 0.2, 0.4, or 0.6M NaCl solution (equivalent to an
osmotic potential of −0.89MPa, −1.77MPa, and −2.66MPa, respectively). He
suggested that priming “Tam Veracruz” pepper seeds for 5days at 23°C in 0.2M
NaCl increased the germination rate at temperatures from 15 to 23°C.A previous
study on pepper (Amjad etal. 2007) showed that halopriming improved seed germi-
nation, seedling emergence, and growth under saline and drought conditions. Khan
etal. (2009a) observed that priming of seeds using NaCl improved seedling vigor
and establishment under salt stress conditions. It has been reported that halopriming
with salt (NaCl) improved the rate of pepper seed germination (Yadav etal. 2011).
In a study comparing hydropriming and halopriming with dehydration treatments
for physiological enhancement of pepper seeds, both hydro- and halopriming treat-
ments with and without dehydration conditions caused increases in normal germi-
nation percentage and germination index of pepper seeds compared with the control
(Sivritepe and Senturk 2011). Maiti etal. (2013) reported that halopriming in 3%
KNO3 solution for 40h at normal room temperature increased speed of emergence,
seedling vigor index, root length and shoot length over hydropriming, and control in
pepper. They also reported that, at eld level, the halopriming treatment increased
yield compared to the control and hydropriming treatments. Dutta etal. (2015) also
reported that pepper seeds primed in 1% KNO3 recorded the highest germination
percentage as compared to nonprimed control.
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
214
12.2.4 Drum Priming
Drum priming is a special seed priming method in which seeds are hydrated to pre-
determine humidity content by placing them inside a horizontal rotating drum.
Drum priming has the capability to accurately hydrate seeds without concern for
spent solution or solid carrier disposal (Rowse 1996). In this novel method, the sup-
ply of water to the seeds is controlled by physical rather than osmotic means or solid
matrix materials, limiting the amount of water introduced to the seed. Specially
designed apparatus enables monitoring of the seed weight, precise regulation of
time, and water amount during hydration process, what ultimately results in an
appropriate and uniform moisture level of the seeds (Warren and Bennett 1997).
The method enables the priming of much larger quantities of seed than previously
been practical and avoids the environmental problems of disposal of the used prim-
ing solution. Unlike other priming methods, it does not require an oxygen enrich-
ment of the atmosphere because of the nature of the process. Drum priming systems
are among the most common methods of seed priming for commercial treatment.
Da Silva etal. (2015) combined drum priming with 24-epibrassinolide (24-EpiBL)
for enhancement of bell pepper seed performance. Several advantages were veried
in the drum priming technique with added 24-EpiBL compared with the traditional
procedure (water alone). Drum priming with 24-epibrassinolide showed positive
effect on germination time and seedling growth of bell pepper.
12.2.5 Solid Matrix Priming
Solid matrix priming (SPM), also known as matriconditioning, is a process in which
seeds are mixed with a solid material and water in known proportions and then
incubated for a given duration at constant temperature. Solid matrix priming utilizes
carriers like vermiculite, diatomaceous earth, or another highly water-absorbent
polymer (Celite® or Micro-cel®) possessing characteristics such as high water
holding capacities, low osmotic potentials, and low bulk density (Taylor etal. 1988,
1998; Khan etal. 1990; Khan 1992; McDonald 2000). In order to improve the con-
trol of imbibition, pure water may be replaced by an osmotic solution, as in osmotic
priming (Khan 1991). After incubation the extraneous solid material is sieved off.
Solid matrix priming is similar to osmotic priming, allowing the seed to imbibe and
attain threshold moisture content and pre-germination metabolic activities but pre-
venting radicle emergence. However, it has the advantages of allowing aeration,
incorporation of biological agents to combat soilborne pathogens, and improved
ease of handling (Taylor etal. 1988; Harman etal. 1989; Paparella etal. 2015). This
technique has been used to improve seed germination and rate in many species
including peppers. Kubik et al. (1988) compared SPM with commercially
osmoprimed tomato and pepper seeds under growth chamber conditions. They used
expanded calcine clay as a solid material. SMP-treated seed performed equally as
well as, or better than, commercially osmoprimed seed. Ilyas etal. (2002) reported
that matriconditioning improved quality and protein level of medium vigor hot
N. Ozbay
215
pepper seed. Kang et al. (2003) compared osmotic with solid matrix priming to
determine the more effective treatment for improving seed germination in pepper
and tomato and reported that solid matrix primed pepper seeds germinate faster than
osmotic primed seeds at all temperatures (15, 20, 25, and 30°C). On the other hand,
they also reported that early growth was not signicantly inuenced by osmotic
priming or SMP treatment of pepper and tomato seeds. Hacisalihoglu and White
(2006) conducted an experiment to determine the optimum duration, temperature,
and ratio (seed/carrier/water) of matriconditioning “long red cayenne” pepper seeds
for improved germination percentage and mean germination time. The results
showed that matriconditioning, which is carried out in 453g glass jars by mixing
seeds, water, and synthetic calcium silicate at 25–30°C for up to 5days, increased
nal germination percentage to 96.7%, compared with 82% in the control seeds.
Furthermore, matriconditioning decreased mean germination time tenfold, com-
pared with a nonprimed control. Pandita etal. (2007) found that solid matrix prim-
ing improved germination of hot pepper seed by 10–16% depending on temperature,
and this effect enhanced when SMP was followed by halopriming and osmoprim-
ing. QianQian etal. (2009) indicated that priming treatment with vermiculite could
improve the vigor of hot pepper seeds and the salt tolerance of hot pepper
seedlings.
12.2.6 Biopriming
Biopriming seed treatment is gaining importance in seed germination and manage-
ment of plant pathogens as another alternative to chemical fungicides in recent
years. The biopriming procedure rst described by Callan etal. (1990) for biologi-
cal control of Pythium preemergence damping-off of sh2 sweet corn integrated
imbibition at an optimal temperature with protection by a biocontrol agent.
Biopriming is a process of biological seed treatment that refers to combination of
seed hydration (physiological aspect of disease control) and inoculation (biological
aspect of disease control) of seed with benecial organisms to protect seed (Rakshit
etal. 2015). The seeds are removed before radicle emergence (Callan etal. 1990).
Because seeds can fail to germinate due to a number of reasons, both biological and
physiological seed treatments used in combination seem to provide the best seed
protection (Bennett etal. 1992). The leakage of seed exudates during biopriming
may supply nutrients and energy for biocontrol agents (Wright etal. 2003). This
favorable environment contributed to colonization and proliferation of biocontrol
agents over the seed surface to facilitate water uptake and nutrients during bioprim-
ing. Biopriming with different benecial microbes may not only enhance seed qual-
ity but also boost seedling vigor and ability to withstand abiotic and biotic stressors
and thus offer an innovative crop protection toll for the sustainable improvement of
crop yield (Rakshit etal. 2015). Biopriming is recently used as an alternative method
for controlling many seed- and soilborne pathogens (Reddy 2012). For example,
combined effect of Pseudomonas uorescens and Trichoderma harzianum as seed
biopriming resulted in signicant growth of pepper seedlings (Kumar etal. 2010).
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
216
In another work on bell pepper, Da Silva and Filho (2012) compared the efciency
of biostimulant priming (10 mL of Stimulate® in 100 mL of distilled water) to
hydropriming in bell pepper seeds as evaluated by SVIS® analysis and recom-
mended vigor tests. They indicated that hydroprimed and biostimulant primed seeds
performed better than control. An investigation was carried out using “PKM 1” chili
(Capsicum annuum L.) seed to standardize biopriming with the biocontrol agents
Trichoderma viride or Pseudomonas uorescens in order to improve seed germina-
tion and seedling vigor (Ananthi et al. 2014). They have suggested that seed
biopriming with 60% (w/v) preparations of T. viride or P. uorescens for 3h or 12h,
respectively, can therefore be adopted to improve seed germination and seedling
vigor in chili. Ilyas etal. (2015) reported that biopriming with the biofungicide and
clove oil 0.06% or 0.1% was an effective seed treatment to improve the vigor and
relative speed of germination while reducing the percentage of Colletotrichum cap-
sici, a seed-borne pathogen causing anthracnose disease in hot pepper seeds (Ilyas
etal. 2015). However, it still provides inconsistent results. Copeland and McDonald
(1995) indicate that, in many cases, the biological treatment of seeds is not persis-
tent in soil, seed, or plant, under natural conditions. Therefore, a deeper research
concerning the methodology of application, doses, storage of coated seeds,
responses from different species to the materials, types of products, and polymers
should be performed to standardize.
12.2.7 Priming withPlant Growth Regulators andOther Organic
Sources
Soaking or treating seeds in optimal concentrations of plant growth regulators
(PGRs) improves germination, stand establishment, growth, and yield of crop
plants under both normal and stress conditions. Carter and Stevens (1998) deter-
mined that when primed with gibberellic acid (GA3), pepper seeds had higher
germination (91%) as compared to nonprimed seeds at high temperature (40°C).
The direct benets of seed priming with PGR were reported by Mendoza etal.
(2002), who demonstrated that that imbibing pepper seeds in 0.1mM salicylic acid
prevented seedling from subsequent chilling-induced damage. Yogananda etal.
(2004) found that bell pepper seeds invigorated with GA3 (200ppm) recorded
higher germination, root and shoot length seedling dry weight, rate of germination,
and seedling vigor indices over control. A laboratory study was performed by Khan
etal. (2012) to explore the benets of seed priming with polyamines on seed ger-
mination and seedling growth of hot pepper. Hot pepper seeds were primed in aer-
ated solution of putrescine, spermine, or spermidine (25, 50, 75, and 100mM) for
48h at 25 ± 2°C.Results showed that polyamine priming resulted in earlier and
synchronized germination via improving nal germination percentage, time to
50% germination, mean germination time, germination energy, germination speed,
and germination index compared with control. Seed performance of various crops
including pepper can also be improved by inclusion of plant growth regulators and
hormones during priming and other pre-sowing treatments (Lee et al. 1998;
N. Ozbay
217
Korkmaz and Korkmaz 2009). For example, priming with PEG and GA4+7 increased
early germination in pepper especially at low temperature (15°C), 3.8 days as
compared 4.7days in nonprimed seeds (Watkins and Cantliffe 1983). Andreoli and
Khan (1999) reported that matriconditioning pepper seeds at 25°C in the presence
of gibberellic acid hastened seed germination and improved stand establishment of
pepper. Korkmaz (2005) reported that inclusion of acetyl salicylic acid and methyl
jasmonate into the priming solution improved low-temperature germination and
emergence of sweet pepper.
Several new compounds are also being tested for their potential use as priming
agents. For example, priming pepper seeds in 3% KNO3 with the presence of
5- aminolevulenic acid (ALA) improved nal germination percentage (FGP) and
germination rate at 15°C compared to nonprimed seeds (Korkmaz and Korkmaz
2009). Ozbay and Susluoglu (2016a) reported that sweet pepper were treated in 3%
KNO3, 2% KH2PO4, and 10% PEG solutions containing 0, 25, 50, and 100mg L-1
prohexadione-calcium (Pro-Ca) in darkness at 25°C for 3days. Priming pepper
seeds, in the presence or absence of plant growth regulator, improved nal germina-
tion percentage, mean time to germination, and germination index, nal emergence
percentage, and mean time to emergence compared to nonprimed seeds. The highest
nal germination percentage and the lowest mean time to germination were obtained
from KH2PO4 + 25mg L-1 Pro-Ca treatment.
Priming with some organic agents could also be used in pepper seeds for physi-
ological enhancement. In an earlier study conducted by Sivritepe and Sivritepe
(2008), possibility of using seaweed extract (as an organic material) in priming
technique was examined in pepper seeds. The seeds (cv. California Wonder) were
subjected to priming treatments performed by the use of 1:1, 1:5, 1:10, 1:25, 1:50,
1:100, 1:250, 1:500, and 1:1000 dilutions of seaweed extract (Maxicrop) and also
H2O, for 1, 2, and 3days at 20°C.They concluded that seaweed extract could be
used as an osmotic agent in organic priming of pepper seeds for physiological
enhancement. They also commented that seaweed extracts caused physiological
enhancements and increases in performance of seeds due to their hormonal compo-
nents such as cytokinins and amino acid contents and also their hygroscopic charac-
teristics. Demirkaya (2010) demonstrated that seaweed extract could be used in
osmotic conditioning treatments of onion seeds as well as pepper seeds. Similarly,
Sivritepe etal. (2015) reported that organic priming with continuously aerated sea-
weed extract (1000ppm) for 2days at 20°C and dehydration treatments gave the
best results in terms of physiological enhancement of pepper seeds. An experiment
was conducted to improve the germination and quality of seedling through seed
priming of bell pepper cv. California Wonder using 16 botanicals and animal by-
products/wastes for 12 and 24h. The best germination and quality of seedlings were
obtained through pre-sowing seed priming treatments of Melia azedarach leaf
extract 10% followed by Eucalyptus leaf extract 10%, garlic clove extract 5%, cow
urine 5%, and cow dung extract 5% at seed soaking duration of 24h (Mehta etal.
2010). Dutta etal. (2015) carried out an experiment to nd out efcacy of organic
(200 g/Kg seed neem leaf powder) and inorganic priming (3% KNO3 and 2%
KH2PO4) on germination and seedling vigor of bird’s eye chili (Capsicum
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
218
frutescens L.) seeds. Among all priming treatments, 1% KNO3 (39.68% higher)
recorded the highest germination percentage as compared to nonprimed control. In
a recent study conducted to investigate effect of organic priming with marigold
herbal tea on seed quality in Aji pepper (Capsicum baccatum var. pendulum Willd.),
the herbal teas obtained from owers of the Tagetes patula and Tagetes erecta spe-
cies were used as an organic priming solution. It has been concluded that organic
priming effectively increased germination, emergence, and the seedling fresh
weight at different maturity levels of pepper (Mavi 2016). On the other hand, Teksan
and Kavak (2016) reported that organic priming treatments with marigold and rose
ower herbal teas did not signicantly affect germination and emergence of pepper
seeds compared to the control. Demirkaya (2016) found that organic priming treat-
ments with seaweed extracts and methyl jasmonate improved the germination per-
centages and reduced the mean germination times of pepper seeds at low
temperatures (15 and 20°C).
12.3 Factors Affecting Priming Success
Priming efciency is affected by many factors such as plant species (even cultivars
within a species), priming technique, priming media, its concentration, water poten-
tial, oxygen availability, priming duration, temperature, seed condition, and storage
conditions (Parera and Cantliffe 1994; Bray 1995). Optimal priming conditions may
vary among cultivars and even seed lots of a given species. Bradford etal. (1990)
indicated that there was a strong interaction between pepper seed lots and priming
response, with the slowly germinating lots exhibiting the greatest benet from prim-
ing. It has been reported that the germination response of two pepper varieties, viz.,
California Wonder and Yolo Wonder, was investigated under the osmotic potential
of −5, −10, and −15 bar created by aqueous solution of PEG 6000 and KNO3.
Variety California Wonder appeared to be more responsive to preconditioning treat-
ments as compared to the variety Yolo Wonder (Thakur etal. 1997). Priming in salt
solutions leads to faster germination than priming in mannitol or PEG (Passam etal.
1989). Similar results were also observed for pepper seeds primed in KNO3 and
KNO3 + K3PO4 solutions (O’Sullivan and Bouw 1984). KN03-primed jalapeno pep-
per seeds resulted in signicantly earlier germination and accelerated vegetative
seedling development, whereas priming in PEG appeared to retard vegetative seed-
ling development (Rivas etal. 1984). In a study conducted by Jones and Sanders
(1987), pepper seeds were soaked in water, a 1% KNO3 + 1% K2HPO4 solution or a
1.5% KNO3 + 1.5% K2HPO4 solution at 21°C for either 72h or 96h. Seeds were
air-dried and germinated at 15°C, 20°C, or 25°C.All soaking treatments hastened
germination and resulted in more uniform germination. Soaking seed in water or a
1% KNO3 + 1% K2HPO4 solution gave similar results and was slightly better in
promoting germination than the 1.5% KNO3 + 1.5% K2HPO4 solution. Hydropriming
(48h) for tomato and sand matrix priming (80% water holding capacity, 3days) for
eggplant and chili were established as best methods of priming treatment capable of
improving seed vigor (Venkatasubramanian and Umarani 2007).
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12.4 Effect ofPriming onSeed Germination andSeedling
Establishment
High yield of the vegetable crops can only be obtained with high stand establish-
ment of seedlings so that they can compete with the environment and can produce
best plantation, ultimately increased yield and quality (Grassbaugh and Bennett
1998; Cantliffe 2003). The nonuniformity of pepper seed germination is one of the
main problems faced by growers, since these seeds usually show low germination
rate and low vigor (Silva etal. 2012). High germination and uniform stand estab-
lishment for chili pepper production are essential to maintaining protable yields.
However, pepper has non-starchy endosperm, and this offered a mechanical barrier
to the growing embryo resulting in poor germination and emergence (Andreoli and
Khan 1999). Nonuniform crop emergence results in plants of variable size and com-
petitive ability. Subsequent management practices may be less effective on elds
with nonuniform emergence. If stand establishment is poor, yields and quality of
once-over machine-harvested crops will be poor. An increasing number of investi-
gators are becoming interested in seed biology with the objective of understanding
and controlling the many aspects of seed germination and seedling establishment
(Bradford 1995). Seed priming is an alternative to achieve this objective. A number
of processes stimulating germination are activated by seed priming and persist after
the dehydration of the seed (Asgedom and Becker 2001). Therefore, upon sowing,
the primed seeds can rapidly imbibe and restore the seed metabolism, resulting in
an increased germination rate, decreased nonuniform germination, and better seed-
ling development (Rowse 1995). Many evidences have shown seed priming could
improve germination and early seedling growth under normal and stress conditions
compared to plants grown from untreated seed (Bradford 1986; Chen etal. 2012).
There are several investigations reporting either no effect or negative or benecial
responses of seed priming for pepper. Ghate and Phatak (1982) reported a signi-
cant decrease in germination rate when pepper seeds were primed with K2HPO4
plus (NH4)2HPO4 solution. It has been also reported that seed priming has not
proved benecial effect for tabasco pepper (Capsicum frutescens L.) eld stand
establishment (Sundstrom etal. 1987). In a study carried out by Jones and Sanders
(1987), pepper seeds were primed in water, a 1% KNO3 + 1% K2HPO4 solution, or
a 1.5% KNO3 + 1.5% K2HPO4 solution at 21°C for either 72h or 96h. Seeds were
air-dried and germinated at 15°C, 20°C, or 25°C.All priming treatments hastened
germination and resulted in more uniform germination. They also reported that
priming seed in water or a 1% KNO3 + 1% K2HPO4 solution gave similar results and
was slightly better in promoting germination than the 1.5% KNO3 + 1.5% K2HPO4
solution. Sundstrom and Edwards (1989) reported that an increased rate of germina-
tion was observed when jalapeno (Capsicum annuum L.) and tabasco (Capsicum
frutescens L.) peppers were primed in a 3.0% or 2.75% KNO3 solution, respec-
tively. In bell pepper, priming of seeds enhanced the rate of germination (Bradford
etal. 1990; Khan et al. 1992). Cooksey etal. (1994) compared non-treated seed,
primed seed, and transplants for effects on stand establishment, plant morphology,
and yield of paprika pepper. They concluded that non-treated seed was satisfactory
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220
for stand establishment, although primed seed had the potential to provide greater
initial stands. They also concluded that transplanting is not recommended for stand
establishment of paprika pepper intended for mechanical harvest. Lanteri et al.
(1994) revealed that priming of pepper seeds in −1.1, −1.3, and −1.5 MPa PEG
solution for 14 days at 25 °C reduced the mean time to germination. Lee et al.
(1997) found that the rate of germination and improvement of seedling stands were
also accelerated as a result of seed priming in pepper. Pepper seeds osmoprimed
with PEG 6000 and KNO3 germinated more rapidly than the control or water pre-
soaked ones. In addition to enhanced germinability, osmopriming treatment was
found to signicantly improve emergence index and vigor index (Thakur et al.
1997). It was also observed that aerated hydration treatment of pepper seeds pro-
duced larger seedlings and better stands in the eld or greenhouse compared to
untreated (Demir and Okcu 2004). It was noticed by Yogananda etal. (2004) that
bell pepper seeds invigorated with GA3 (200ppm) or KNO3 (1.0%) recorded higher
germination, root and shoot length, seeding dry weight, rate of germination, and
seedling vigor index over control. Korkmaz (2005) for sweet pepper reported that
priming treatments generally improved the germination synchrony. In capsicum,
Pandita etal. (2007) observed that osmo- and solid matrix priming improved seed
germination over nonprimed seeds at 15°C, 20°C, and 25°C temperatures. They
also observed that priming also reduced mean days to germination signicantly over
control. It was reported by QianQian etal. (2009) that after priming treatment of the
seeds with vermiculite, the germination rate, the germination energy, the germina-
tion index and vigor index of hot pepper seeds, and the fresh and dry weights of
seedlings were signicantly higher than those of control. Yadav etal. (2011) noticed
that germination percentage of primed pepper seeds was increased to as compared
to control and also tolerated cold and salt stress for 10days with 100% survival,
whereas control seedling could not survive. Ameri etal. (2011) conducted a labora-
tory experiment on seed priming of pepper “California Wonder” with 1% NaCl, 1%
CaCl2, 3% KNO3, 3% FeSO4, and control. All treatments resulted in a higher germi-
nation rates compared to the control. They also indicated that seed priming with
FeSO4 was the best treatment resulting in the maximum radicle dry weight, germi-
nation percentage, and germination rate with values of 0.126%, 70%, and 5.07%,
respectively, while in the control values were 0.063%, 36.81%, and 0.83%, respec-
tively. In a work investigating effects of osmotic conditioning and humidication
applications on emergence percentage and mean emergence time of pepper seeds
(Demirkaya 2012), the seeds were osmoprimed with PEG-6000 and −1.0MPa for
1, 2, and 3days and hydroprimed for 1, 2, and 3days. The results of the study indi-
cated that humidication and priming applications with PEG-6000 positively
affected mean emergence time and emergence percentages in pepper seeds.
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12.5 Effect ofPriming onPerformances ofPepper Seeds
atLow Temperature
Low temperature is one of the most important environmental factors affecting pep-
per seed germination or seedling emergence. Most of the pepper cultivars are
extremely sensitive to chilling stress particularly during emergence and early stages
of seedling development. However, different varieties of the same species can toler-
ate low temperature differently. Peppers are very sensitive to germination tempera-
tures, tolerating 16°C on the lower end, with an optimum range between 18 and
35°C and a mean optimum of 29.5°C (Nonnecke 1996). The rate of germination
and emergence of pepper seeds is markedly reduced at a temperature ranging from
15 to 20 °C. Pepper plants experience chilling injury with prolonged temps of
0–10° C.Methods to bypass the slow germination of pepper seed at low tempera-
tures have been studied by researchers. Seed priming is an effective and practical
technique that enables the seed to germinate and emerge faster even at suboptimal
temperatures and also known to reduce the imbibitional damage associated with
planting seeds in cold soils (Bennett and Waters 1987; Jisha etal. 2013). The effect
of priming on low-temperature performance of pepper seeds has ranged from no
improvement to some advancement in germination and emergence percentages and
rates. For example, imbibition at 30°C for 48h in water or in aerated KNO3 solu-
tions for 6 or 8days enhanced the germination at 15 °C when the seeds were not
redried after treatment (Sachs etal. 1980). It has been reported that eld emergence
percentages of pepper at 20 °C generally were unaffected by priming (Bradford
etal. 1990). Carter (1994) initiated to examine the effects of NaCl priming on total
germination and the rate of germination under suboptimal temperature conditions.
Seeds of “Tam Veracruz” chili were primed at 23°C for 5days in double-distilled
H2O or 0.2, 0.4, or 0.6 M NaCl solution (equivalent to an osmotic potential of
−0.89MPa, −1.77MPa, and −2.66MPa, respectively). He suggested that priming
“Tam Veracruz” chili seed for 5days at 23°C in 0.2M NaCl increased the germina-
tion rate at temperatures from 15 to 23°C.Matriconditioning pepper seeds at 15°C
for 7days or at 25°C for 4days reduced the time needed for germination on lter
paper at 15°C, and conditioning at 25°C was more effective than conditioning at
15°C in reducing the germination time. The priming treatment also improved the
performance of pepper seeds in early eld plantings at suboptimal temperatures
(averaged over 10days after planting) ranging from 12 to 18°C (Khan etal. 1995).
Increasing evidence suggests that benzoic acid derivatives such as salicylic acid
(SA) or ASA regulate stress tolerance in plants (Lopez-Delgado et al. 1998).
Mendoza etal. (2002) reported that priming pepper seeds in 0.1mM SA prevented
seedlings from subsequent chilling-induced damage. The incorporation of plant
growth regulators into the priming solution has been shown to be effective to
enhance germination and seedling emergence of crops under adverse condition. For
example, incorporation of acetyl salicylic acid (ASA) or methyl jasmonate (MeJA)
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
222
into the priming solution improved sweet pepper germination and emergence at low
temperature (Korkmaz 2005). Priming sweet pepper seeds in KNO3 supplemented
with 0.1mM acetyl salicylic acid resulted in 91% germination and 85% emergence
at 15 °C which are signicantly higher than the germination and emergence of
nonprimed and seeds primed in KNO3 only (Korkmaz 2005). In the same study, it
has been also reported that the primed seeds stored for 1month at 4°C still exhib-
ited improved germination performance at 15°C.Similarly, Korkmaz and Korkmaz
(2009) indicate that priming seeds in 25ppm and 50ppm ALA (5-aminolevulinic
acid) incorporated into the KNO3 solution improved low-temperature performance
of red pepper seeds. They also indicated that primed seeds stored for 1month at
4°C or 25°C still exhibited improved germination and emergence performance at
15 °C. Chemical seed priming treatments imparted tolerance to subsequent cold
(4°C) or salt stress (NaCl, 200 mM) exposure at later growth stages. Among the
treatments, seed priming with thiourea (TU, 1.3mM) was effective in imparting
cold as well as salt stress tolerance (Yadav etal. 2011). In a recent work carried out
by Sharma etal. (2015), the effect of seed priming using PEG-6000, gibberellic
acid, KH2PO4, Na2HPO4, distilled water, and cow urine on seedling vigor of bell
pepper seeds was studied. The seeds were primed at 20°C for 24 and 48h. They
reported that although all the priming treatments signicantly improved seed per-
formance over control, seed priming with 100ppm GA3 for 48h was more effective
in improving seed performances under low temperature than the other treatments.
The addition of prohexadione-Ca into the priming solution signicantly improved
sweet pepper germination and emergence at low temperatures compared to unprimed
seeds and the seeds primed in KNO3, KH2PO4, and PEG only (Ozbay and Susluoglu
2016a). Samarah etal. (2016) compared hydropriming and natural compounds that
could be used as seed treatments to increase seed germination rate and improve cold
tolerance of bell pepper. They reported that treatments with nanochitin, chitosan,
acetic acid, or hydropriming improved low-temperature germination in soil by
17–39% compared with untreated seeds, being hydropriming or nanochitin more
effective in reducing mean time to germination than chitosan or acetic acid treat-
ments. Result of the study conducted by Korkmaz etal. (2017) revealed that seed
application of 1μM or 5μM melatonin signicantly improved pepper seed germi-
nation and seedling emergence at chilling temperatures compared to seeds not
treated with melatonin.
12.6 Effect ofPriming onPerformances ofPepper Seeds
atHigh Temperature
High temperature is also one of the most important environmental factors affecting
seed germination and seedling emergence of many vegetable crops including pepper.
Pepper seeds for fall production are sown when summer greenhouse temperatures
can reach 40–45°C (Vavrina 1994). This range is far above the optimum temperature
(29 °C) for pepper germination (Maynard and Hochmuth 1997). At supraoptimal
temperatures, pepper seeds may enter into the state of thermoinhibition. The inability
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to germinate at higher than optimal temperatures is attributed to a condition called
thermoinhibition or thermodormancy (Carter and Varina 2001). Most of the pepper
cultivars are extremely sensitive to high temperature during germination and emer-
gence and also later stages of plant development such as owering and fruit set
(Aloni etal. 2001). However, different varieties of the same species can tolerate high
temperature differently as in low temperature mentioned previously. Cultivar differ-
ences in nal germination pronounced at 35°C.Therefore, growers should consider
this, when choosing cultivars for fall transplants. However, no cultivar germinated
well at temperatures higher than 35°C (Carter 2000). Priming can also help alleviate
high-temperature inhibition of germination and improve seedling emergence of pep-
per and other species (Ellis and Butcher 1988; Cantliffe 1991). For example, pepper
seed germination was improved under supraoptimal conditions if the seeds were rst
soaked in solutions of ethephon or gibberellic acid (GA3) (Carter and Stevens 1998).
Nascimento etal. (2011) reported that osmoconditioning improves the physiological
quality of hot pepper seeds by improving germination at high temperature (35°C). In
a study conducted by Silva etal. (2012), hot pepper “Mari” seeds were osmocondi-
tioned in aerated solution of polyethylene glycol (PEG 6000) for 7days and sub-
jected to germination test at 15, 20, 25, 30, and 35°C. The total seed germination
decreased with increasing temperature. The osmotic conditioning was effective in
improving seed germination at all temperatures, especially at high temperatures.
Ozbay and Susluoglu (2016b) reported that priming pepper seeds in the presence or
absence of plant growth regulators improved nal germination and emergence per-
centage, mean emergence time, emergence index, and emergence rate (E50) at high
temperature (35°C) compared to control seeds.
12.7 Priming toOvercome Salt Stress
Among the abiotic stresses, salinity is a major limiting factor in the crop productiv-
ity all over the world. Almost 20% of cultivated area of the world and half of the
world’s irrigated lands are stressed by the salinity (Chinnusamy etal. 2005). Soil
salinity, if not properly managed, often results in poor stand establishment, reduced
plant growth, and reduced yield of many horticultural crops such as peppers (Flynn
etal. 2003; Niu etal. 2010). When exposed to high salt concentration, growth of
pepper, which is not a salt-tolerant crop, can be negatively affected, and yield and
fruit quality are diminished (Ibn Maaouia-Houimli etal. 2011). Salinity may reduce
crop yield by upsetting water and nutritional balance of the plant (Khan etal. 2007).
The salinity delays or prevents the seed germination through various factors, such
as a reduction in water availability by high osmotic potential and toxicity of Na and
Cl ions, changes in the mobilization of stored reserves, and affecting the structural
organization of proteins (Kumar 1995; Ibrahim 2016). Salt concentrations affected
percentage and rate of germination as well as the opening of cotyledonary leaves in
Capsicum annuum L. cv. California Wonder. The 100mM or higher salt concentra-
tions reduced the rate of radicle protrusion signicantly (Yadav et al. 2011).
Similarly, Chartzoulakis and Klapaki (2000) reported the signicant reduction in
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
224
the germination as well as growth of two bell pepper hybrids on exposure to 100 and
150mM NaCl. Furthermore, pepper yield is reduced up to 14% for every increase
in unit of salinity above its threshold (Rhoades etal. 1992). To alleviate this prob-
lem, a number of studies were conducted with the aim of removing the inhibitory
effect of salt stress on plant growth. The benecial effect of seed priming has been
used for improving germination, emergence, and stand establishment of peppers
under salt conditions (Hassen etal. 2014). Plants derived from primed seed have a
higher adaptation capacity to salinity originating in the osmoregulation process
(Levitt 1980). In studies conducted by Smith and Cobb (1991a), Amjad etal. (2007),
and Yadav etal. (2011) with pepper, it was concluded that seed priming improves
seed germination, seedling emergence, and growth under saline conditions. Priming
of seeds with NaCl was found to be effective in alleviating the adverse effects of salt
stress on pepper plants. Khan etal. (2009a) observed that priming of seeds using
1mM NaCl improved seedling vigor and establishment under salt stress conditions.
It has been reported that hormonal priming with salicylic acid (0.8mM) and acetyl-
salicylic acid (0.2 mM) showed signicantly better results over the control by
improvement in time taken to 50% emergence, nal emergence percentage, root and
shoot length, seedling fresh and dry weight, and seedling vigor under normal as well
as saline conditions. It has also been reported that acetylsalicylic acid exhibited
superiority over salicylic acid (Khan etal. 2009b). Pepper seeds treated with 10mM
glyciebetaine (GB), an organic osmolyte accumulated in variety of plants in
response to abiotic stress, for 24h in darkness improved germination and synchrony
of germination under salt stress. Glyciebetaine decreases the melonoldehyde (MDA)
and increases the proline content and SOD activity in seeds. The enhanced tolerance
to salt stress may be due to reduced lipid peroxidation and elevated SOD enzyme
activity (Korkmaz and Sirikci 2011). Osmopriming with CaCl2, KCl, and NaCl
improved germination rate, chlorophyll content, proline, and protein accumulation
in pepper under salt stress conditions (Hassen etal. 2014). Hassen etal. (2017) also
conducted a study to assess the effects of priming with concentrations of KCl, NaCl,
and CaCl2 on morphological and biochemical parameters of the pepper cvs. Beldi,
Baklouti, and Anaheim chili. They reported that seedlings developed from primed
seed had improved biomass, water content, carotenoid content, soluble sugar, poly-
phenols, and soluble proteins at salt concentrations of 6g·L−1.
12.8 Reversal ofSeed Deterioration by Priming
Another area of practical concern of priming is the interaction between priming
treatments and seed deterioration. Seed deterioration may be dened as the loss of
seed viability and vigor due aging effects and adverse environmental factors par-
ticularly higher temperature, relative air humidity, and oxygen/carbon dioxide ratio
(McDonald 1999; Jyoti and Malik 2013). Seed deterioration is associated with vari-
ous cellular, metabolic, and chemical alterations including lipid peroxidation, mem-
brane disruption, DNA damage, impairment of RNA, and protein synthesis and
causes several detrimental effects on seed (Jyoti and Malik 2013). Seed storage
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causes a decrease in the protein content which may be related to oxidation of the
amino acids due to the increase in the respiratory activity and advance in the dete-
rioration process of the stored seeds (Manonmani etal. 2014). Poor storage condi-
tions may accelerate seed deterioration in primed and unprimed seeds (Georghiou
etal. 1987). It is a serious problem particularly in developing countries where seeds
are stored in places usually without a proper control of air humidity, temperature,
and O2/CO2 concentration (Khan etal. 2016). As seed deterioration increases, seed
performance progressively decreases. Plants that have originated from deteriorate
seed can also reduce growth rate (Kapoor etal. 2010). Pepper is a warm climate
crop, and seeds are sensitive to storage conditions and lose germination potential
within a short period of time, particularly under adverse storage conditions (Priestly
1986). Decline in germination potential due to seed aging may result in low seedling
emergence and stand establishment (Ermis etal. 2016). Accelerated aging of sweet
pepper seeds at 42°C and 100% R.H. for 0, 5, 10, 15, 20, 25, and 30days resulted
in decreased germinability in terms of radical emergence (%) that was also differed
signicantly among different aging duration (Kaewnaree etal. 2011). According to
Li etal. (2005), the main mechanism for aging of pepper seeds is associated with
increased peroxidation of lipid membranes. The aging of pepper seeds, during long-
term storage, deteriorated their vital status which was expressed in changes in their
moisture content, decreasing of their sowing qualities, and development of weaker
seedlings with higher water content (Panayotov and Aladjadjiyan 2014). Priming
can reverse some of the aging-induced deteriorative factors and thus improve seed
performance (Taylor etal. 1998). The benecial effects of priming are associated
with the repair and building up of nucleic acid, increased synthesis of proteins, as
well as the repair of both mitochondria and membranes (McDonald 1999, 2000).
Under invigoration, metabolic repair processes in deteriorated seeds occur before
onset of seed germination process (Srinivasan etal. 2009). Many seed priming treat-
ments have been used to reduce the damage of aging and invigorate their perfor-
mance in many crops including pepper (Farooq etal. 2009; Ermis etal. 2016). In
pepper, osmoconditioning was found to be also an effective treatment for protecting
seeds stored under high temperatures by increasing longevity of seeds. However,
osmoconditioning after storage did not seem to have any signicant effect on seed
viability, though it enhanced the germination rate (Georghiou et al. 1987).
Osmoconditioning of controlled deteriorated sweet pepper seeds exerted various
effects on seed germination depending on deterioration rate (Lanteri etal. 1996,
1997). Lanteri etal. (1996) reported that effects of aging pepper seeds may be partly
reversed by priming and part of priming effects is related to the induction of nuclear
replication. In a study conducted by Siri etal. (2013), sweet pepper seeds were arti-
cially aged by exposing to high temperature (42°C) and high humidity (100%
relative humidity) and then primed with PEG 6000 solution with the osmotic poten-
tial of −1.5MPa for 6days. They concluded that seed germination was improved in
primed seeds due to accumulation of antioxidants and the improvement of cell
membrane integrity. Ermiş et al. (2016) indicate that priming is more useful for
enhancing germination of low-quality seed lots than higher-quality ones which indi-
cates that repair of aging is one of the primary advantages of the priming treatments.
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
226
While in general it is clear that the best commercial potential for priming is the
enhancement of performance of seeds of the highest quality (Perkins-Veazie and
Cantliffe 1984), such repair treatments may be appropriate in the cases of valuable
deteriorated stock or rare genetic material.
12.9 Storage Life ofPrimed Seeds
It would be highly advantageous if the promotive effects of priming on the germina-
tion of seeds were retained after drying and storage. In general, storing primed seeds
reduces germination or seedling emergence, and there are conicting results on the
effect of storage life of primed seeds. As far as it concerns the storage of pepper
seeds following priming treatments, Bruggink etal. (1999) reported that for pepper
seeds, the desired longevity was obtained by keeping the seeds, after a priming
treatment, under a mild water and/or temperature stress for a period of several hours
to days. O’Sullivan and Bouw (1984) reported that a promotive effect on the germi-
nation of pepper seeds, when primed in 15% salt solution, was retained for 2days
after treatment. Perl and Feder (1981) showed a retention of seed vigor and seedling
development of pepper seeds for 2months following priming. Aljaro and Wyneken
(1985) reported the effects of priming of “Y010 Y” pepper seeds to be apparent for
up to 140days prior to sowing. In pepper, osmoconditioning was found to be also
an effective treatment for protecting seeds stored under high temperatures
(Georghiou etal. 1987). Thanos etal. (1989) reported that primed sweet pepper
seeds retained their improved vigor after a storage period of 6months at 5–8°C.It
was determined that primed pepper seeds with PEG could be packaged in original
packages and stored for 12months under controlled conditions without any loss of
effect on emergence time (Cetin and Duman 2005). They also reported that air- and
moisture-proof aluminum materials were the best seed storage packaging material
for the primed seeds. Korkmaz (2005) reported that 1month storage at 4–8°C did
not reduce the performance of pepper seeds that were primed in the presence of sali-
cylic acid. Korkmaz and Korkmaz (2009) indicate that red pepper seeds primed in
25ppm and 50ppm ALA (5-aminolevulinic acid) incorporated into the KNO3 solu-
tion can be stored for 1month at 4°C or 25°C and still exhibit improved germina-
tion and emergence performance at 15°C.On the contrary, in other experiments, the
promotive effect of priming was reduced rapidly after drying of the seeds (Sachs
etal. 1980; O‘Sullivan and Bouw 1984). The response of primed seeds to storage is
species and variety dependent. Therefore, it is important that for the benecial
effects of priming to be retained, new techniques and markers have to be used to
monitor the priming progress.
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12.10 Effect ofPriming onPhysiological andBiochemical
Changes inPepper Seeds
The positive effects of priming on the germination performance of many species
might be attributed to the induction of physiological and biochemical changes such
as DNA replication, increased RNA and protein synthesis, greater ATP availability,
increased respiratory activity, faster embryo growth, reduced leakage of metabo-
lites, and decrease in lipid peroxidation and increased in the antioxidant activities.
12.10.1 Seed Priming andCell Cycle Regulation
Some of the hypotheses proposing explanation for priming-induced improvement
of seed performance are based on its effect on DNA in relation to activation of DNA
repair mechanisms, synchronization of the cell cycle in G2, and preparation to cell
division (Lutts etal. 2016). For example, Stofella etal. (1992) indicate that the
increase in root/shoot ratio with hydropriming treatments may be due to the fact that
priming induced nuclear replication in root tips of fresh seeds. By incorporating
DNA-specic uorescent dyes, nuclear DNA contents expressed as C values have
been quantied in pepper (Lanteri etal. 1993) seeds during germination and prim-
ing. Lanteri etal. (1994) reported that priming of pepper and tomato seeds in −1.1,
−1.3, and −1.5MPa PEG solution for 14days at 25 °C reduced the mean time to
germination. Lanteri etal. (1996) reported that effects of aging pepper seeds may be
partly reversed by priming and part of priming effects is related to the induction of
nuclear replication. It has been reported that priming of seed promotes germination
by repair of the damaged protein, RNA, and DNA (Koehler etal. 1997). An induc-
tion of 4C signals was also found after priming, indicating that during priming the
cells of the embryonic root tip had replicated their DNA and arrested at the G2
phase of the cell cycle. Flow cytometric determination of nuclear DNA contents in
embryos of dry, fully matured pepper seeds revealed only 2C signals. Therefore,
pepper belongs to those species in which the quiescent embryo arrests nuclear divi-
sion in the presynthetic G1 phase. 4C nuclei appear 1–2days after imbibition in
water, while radicle emergence starts 2days later; at this time the proportion of the
4C nuclei is over 50% (Lanteri etal. 1992, 1993; Saracco etal. 1995). Thus, DNA
replication precedes pepper seedling growth, and the 4C/2C ratio can be used to
predict seedling performance of pepper (Sliwinska 2009). Using ow cytometry,
Lanteri et al. (1997) observed that priming treatments in PEG solutions might
induce DNA replication in the embryo root tips of pepper seeds. Lanteri etal. (1998)
have also reported that the improvement of performance in pepper seeds after con-
ditioning osmotic activity has been correlated with macromolecular repair that
occur during the treatment. Changes in nuclear replication stages upon priming
have been studied by ow cytometry on pepper seeds. The author observed that
signicant correlations were found between the frequency of priming-induced
nuclear replication and the improvement of pepper seed vigor, as measured by the
reduction in mean germination time (Sliwinska 2009). Flow cytometric data
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
228
published by Varier et al. (2010) also reveal that the improvement of germination
associated with priming is accompanied by increase in 4C nuclear DNA.This indi-
cates that priming enhances DNA replication allowing the advancement of the cell
cycle from G1 to G2 phase.
12.10.2 Effect ofPriming onSeed Ultrastructure
The seeds of some species including peppers are prevented from completing germi-
nation because the embryo is constrained by seed coats and surrounding structures
(Watkins and Cantliffe 1983; Bradford 1995; Bewley 1997a). After imbibition of
the seeds, however, the endosperm tissue enclosing the embryo restrains the germi-
nation process acting as a physical barrier, which restricts radicle emergence (Bino
etal. 1998; Lutts etal. 2016). It is necessary to reduce the resistance of these enclos-
ing structures for germination to be completed. Therefore, weakening of the endo-
sperm, particularly in the micropylar region adjacent to the radicle, is a prerequisite
for the completion of germination (Gong etal. 2005). Priming may also contribute
to rapid seed germination by reducing the mechanical restraint of endosperm on
developing embryo (Mayer and Mayber 1989). In studies with pepper seeds,
Watkins and Cantliffe (1983) and Watkins etal. (1985) found that the mechanical
constraint by endosperm in the tip region of the radicle contributed to slow germina-
tion rate and that the weakening of the endosperm by externally applied gibberellic
acid (GA4+7) occurred prior to germination. The reserves in endosperm are degraded
by specic enzymes after the initiation of germination to facilitate the developing
seedling until photosynthesis is initiated (Bewley and Black 1994; Homrichhausen
etal. 2003). In most cases, endo-beta-mannanase degrades endosperm cell walls in
endospermic seeds and can be detected during germination in the seeds (Bewley
1997b; Cantliffe 2003). Andreoli and Khan (1999) indicate that the matricondition-
ing seed treatment provides a means to efciently digest the endosperm cell by
GA-induced enzymes and reduce the mechanical restraints of endosperm thus pro-
viding energy to start and sustain embryo growth of tomato and pepper. Lanteri
etal. (2000) investigated the expression of β-tubulin in the root tips of pepper seeds
as a complementary marker for priming de novo synthesis of β-tubulins in response
to priming was observed prior to DNA replication after osmopriming for different
durations at two water potentials. Ethylene also directly inuences germination
speed and percentage. Increase in ethylene production during priming may promote
endo-β-mannase activity facilitating endosperm weakening and post-priming ger-
mination (Chen and Arora 2013). Priming was reported to modify the kinetics of
ethylene synthesis from its precursor 1-aminocyclopropane-1-carboxylic acid
(ACC) (Wu et al. 2014). It has been reported that the primed pepper seeds had
higher ACC level, greater ACC-oxidase activity, and greater embryo growth poten-
tial than the nonprime seeds (Khan etal. 1995).
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229
12.10.3 Effect ofPriming onReserve Mobilization
It is proposed that germination-related processes such as respiration, energy
metabolism, and early reserve mobilization can also occur during priming (Varier
etal. 2010). Higher respiratory activity is required to cover energy pool for speed
up germination. Increased respiratory activity has been reportedly associated with
pre- sowing treatments. For example, Halpin-Ingham and Sundstrom (1992) indi-
cate that priming increases respiratory activity of pepper seeds. They also reported
that seed priming generally reduced the oxygen-time constant and increased the
standard deviation of germination responses. During seed germination, storage
proteins, which provide a source of reduced nitrogen, and inorganic minerals need
to be mobilized to support seedling growth (Lutts etal. 2016). Pepper seeds were
examined during priming to determine if seed treatments which accelerate the rate
of germination could be correlated with specic physiological changes within the
seeds. Smith and Comb (1991b) found that soluble protein content increased to
109% and 120% in pepper seeds primed in −0.90 and −1.35MPa NaCl solutions,
respectively, after 12days of priming and also revealed that there was no signi-
cant difference in the soluble protein content between two priming treatments.
Khan (1992) found that two amino acids were incorporated in proteins during the
rst 24h of imbibition of sweet pepper seeds in PEG solutions. QianQian etal.
(2009) reported that the priming treatment with vermiculite signicantly resulted
in signicantly higher contents of soluble protein in the pepper seedlings under
NaCl stress. Hassen etal. (2017) reported that pepper seedlings developed from
primed seed had improved soluble proteins at salt concentrations of 6g·L−1. In
contrary, in a study conducted to determine the relationship between physical and
metabolic changes and germination/emergence time and rates both under optimum
and stress conditions during osmotic conditioning process in pepper seeds,
Ozpercin etal. (2005) reported that no signicant changes were found in total pro-
tein levels in primed pepper seeds. Osmopriming induced accumulation of stress
proteins, such as late embryogenesis abundant (LEA) proteins and heat shock pro-
teins (HSP) (Gallardo etal. 2002). As LEA proteins accumulate at a high level in
response to cell/tissue dehydration, they may contribute to acquisition of tolerance
to drought and related stresses such as osmotic, salt, and cold stress (Lutts etal.
2016). Cortez-Baheza et al. (2007) have shown that several late embryogenesis
abundant (lea) genes are strongly induced in pepper seeds when osmoprimed with
PEG incorporated with GA3. A new LEA protein of 73 amino acids (Calea 73
gene) was highly induced in osmoprimed treatments in which KNO3 was used in
combination with PEG on C. annuum cv. Caballero seeds. Besides being induced
by PEG + GA3, the Calea 73 gene was also stimulated by PEG + KNO3, which
indicates that this gene is expressed during osmopriming regardless of the osmotic
solution used (Cortez-Baheza etal. 2008).
12 Studies onSeed Priming inPepper (Capsicum annuum L.)
230
12.10.4 Effect ofPriming onOsmoprotectants
The osmoprotectants are compounds produced in plants during osmotic stress con-
dition. Osmoprotectants are chemically small, electrically neutral molecules, which
play important roles in the protection and stabilization of proteins and membranes
against abiotic stresses without disrupting plant metabolism (Yancey 1994).
Osmoprotectants or compatible solutes include proline, mannitol, D-ononitol, tre-
halose, sucrose, fructane, glycine betaine, and polyamines (Khan etal. 2015; Lutts
etal. 2016). Accumulation of osmoprotectants such as proline and sugars has been
reported in various plant species including peppers during a wide range of abiotic
stresses. Proline is a proteogenic amino acid and accumulates both under stress and
non-stress conditions as a benecial solute in plants (Kavi Kishor etal. 2015). The
main function of proline in plants is to restrain the osmotic effects by stabilizing
protein structures and scavenging free radicals (Smirnof and Cumbes 1989;
Biedermannova etal. 2008). Accumulation of sugars in various plant species is
related to high tolerance to stress conditions (Bohnert and Jensen 1996). Sugars
protect membranes and enzyme complexes from reactive oxygen species mainly by
interacting with enzymes of the glutathione–ascorbate cycle (Khan et al. 2015).
There are several reports showing that the osmoprotectant content increases after
seed priming. For example, the priming pepper seeds with vermiculite signicantly
increased content of free proline and soluble sugar in the seedlings under NaCl
stress (QianQian et al. (2009). Korkmaz and Şirikçi (2011) reported that pepper
seed treated with glyciebetaine exhibited an enhancement in proline content under
saline conditions. Hassen etal. (2014) showed that priming pepper seeds with NaCl,
KCl, and CaCl2 caused a marked increase in proline accumulation. Seedlings devel-
oped from primed pepper seeds had improved biomass, water content, carotenoid
content, soluble sugar, and polyphenols at salt concentrations of 6g·L−1 (Hassen
etal. 2017).
12.10.5 Effect ofPriming onManagement ofOxidative Status
Management of oxidative status is also an important part of primed seed physiology
(McDonald 2000; Kibinza etal. 2011). The priming could activate the response of
the antioxidant system, becoming the primed seeds more prepared for possible
stresses. During seed imbibition and early stages of germination, reactive oxygen
species (ROS) production occurs mainly through respiratory activities of mitochon-
dria, activities of β-oxidation pathways and enzymes such as NADPH oxidases,
extracellular peroxidases, and oxalate oxidases (Wojtyla etal. 2016). Antioxidants,
by scavenging the excessive ROS during early imbibition, play an essential role in
ensuring successful germination, especially under stress conditions (Bailly 2004).
An increased activity of antioxidant enzymes like ascorbate peroxidases (APX),
catalase (CAT), peroxidase (POX/POD), glutathione reductase (GR), and superox-
ide dismutase (SOD) as a consequence of seed priming has been reported in various
crops including peppers. For example, the priming treatment with vermiculite
N. Ozbay
231
signicantly improved the activities of superoxide dismutase (SOD) and peroxidase
(POD) in the leaves of seedlings and signicantly decreased the rate of producing
superoxide anion (O2-) and the content of malondialdehyde (MDA), resulting in
lower membrane lipid peroxidation in the seedlings under NaCl stress (QianQian
etal. 2009). Kaya etal. (2010) reported that the most important effects of priming
on enzymatic activities of pepper seeds were recorded in catalase which increased
remarkably with priming. They further stated that, even though not the same extent,
priming also increased ascorbate peroxidase and superoxide dismutase activities
(Kaya etal. 2010). Korkmaz etal. (2010) reported that 5-aminolevulinic acid (ALA)
pretreatment increased relative water content, stomatal conductance, and superox-
ide dismutase (SOD) enzyme activity and reduced membrane permeability in pep-
per. Pepper seeds treated with glyciebetaine exhibited a signicant decrease in
MDA content and enhancement in SOD enzyme activity under saline conditions
(Korkmaz and Sirikci 2011). In a study focusing on the effect of priming process on
quality and biochemical changes in sweet pepper seed, Siri etal. (2013) reported
that osmopriming with −1.5MPa PEG-6000 for 6days of articially aged seeds of
sweet pepper (42°C and 100% relative humidity) resulted in an improved germina-
tion with decreased levels of malondialdehyde (MDA) and total peroxide concen-
tration. They further stated that accumulation of total antioxidant activity (TAA),
total ascorbate, dehydroascorbate, and catalase (CAT) activity in primed seeds
enhanced the defense mechanism in protecting the cell membrane damage from
reactive oxygen species. Da Silva etal. (2015) showed that priming improved super-
oxide dismutase (SOD), catalase (CAT), and peroxidase (POX) activities.
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